Carbon isotope composition in relation to leaf gas exchange and environmental conditions in Hawaiian Metrosideros polymorpha populations

Summary Carbon isotope composition, photosynthetic gas exchange, and nitrogen content were measured in leaves of three varieties of Metrosideros polymorpha growing in sites presenting a variety of precipitation, temperature and edaphic regimes. The eight populations studied could be divided into two groups on the basis of their mean foliar ¹³C values, one group consisting of three populations with mean ¹³C values ca.-26 and another group with ¹³C values ca.-28. Less negative ¹³C values appeared to be associated with reduced physiological availability of soil moisture resulting from hypoxic conditions at a poorly drained high elevation bog site and from low precipitation at a welldrained, low elevation leeward site. Gas exchange measurements indicated that foliar ¹³C and intrinsic wateruse efficiency were positively correlated. Maximum photosynthetic rates were nearly constant while maximum stomatal conductance varied substantially in individuals with foliar ¹³C ranging from-29 to-24. In contrast with the patterns of ¹³C observed, leaf nitrogen content appeared to be genetically determined and independent of site characteristics. Photosynthetic nitrogenuse efficiency was nearly constant over the range of ¹³C observed, suggesting that a compromise between intrinsic water- and N-use efficiency did not occur. In one population variations in foliar ¹³C and gas exchange with leaf cohort age, caused the ratio of intercellular to atmospheric partial pressure of CO₂ predicted from gas exchange and that calculated from ¹³C to be in close agreement only in the two youngest cohorts of fully expanded leaves. The results indicated that with suitable precautions concerning measurement protocol, foliar ¹³C and gas exchange measurements were reliable indicators of potential resource use efficiency by M. polymorpha along environmental gradients.     

Morphological and physiological responses of two coffee progenies to soil water availability

Drought is a major environmental constraint affecting growth and production of coffee. The effects of water supply on growth, biomass allocation, water relations, and gas exchange in two coffee progenies representing drought-tolerant (Siriema) and drought-sensitive (Catucaí) genotypes were compared. They were grown in 12-L pots until 4-months old, when they were submitted to two watering treatments for 60 d: plants receiving either 100% transpired water (control plants) or a fraction (about 40%) of the amount of water transpired by control plants (drought-stressed plants). Under control conditions, Siriema grew faster than Catucaí. Regardless of the watering regimes and progenies, relative growth rate (RGR) was positively correlated both with net assimilation rate (NAR) and long-term water-use efficiency (WUE), but not with differences in biomass allocation. Both progenies responded to drought stress through (i) similar decreases in both RGR and NAR with marginal, if any, changes in allocation; (ii) decreases in leaf water potential, which occurred to a greater extent in Catucaí than in Siriema, even though they have showed similar abilities to adjust osmotically and elastically; (iii) similar reductions in net photosynthesis due mainly to nonstomatal factors; and (iv) decreases in transpiration rate coupled with increased long-term WUE. However, the lower transpiration rate and the higher long-term WUE as found in Siriema relative to Catucaí under control conditions persisted under drought conditions. Overall, the major differences between these progenies were largely associated with differences in plant water use, which was likely related to the improved water status of Siriema. The possible implications of selecting coffee genotypes for high WUE are discussed.     

Homeostatic gas-exchange parameters inferred from 13C/12C in tree rings of conifers

The CO₂ concentration of the atmosphere has increased by almost 30% in the past two centuries, with most of the increase (>5 Pa) during the past 60 years. Controlled environment studies of crop plants dependent on the C₃ photosynthetic pathway indicate that an increase of this magnitude would enhance net photosynthesis, reduce stomatal conductance, and increase the difference in CO₂ concentration across the stomata, i.e., CO₂ concentration outside the leaf to that within (c _a-c _i). Here we report evidence, based on stable isotope composition of tree rings from three species of field-grown, native conifer trees, that the trees have indeed responded. However, rather than increasing c _a-c _i, intercellular CO₂ concentrations have shifted upward to match the rise in atmospheric concentrations, holding c _a-c _i constant. No differences were detected among Douglas-fir (Pseudotsuga menziesii), ponderosa pine (Pinus ponderosa), or western white pine (Pinus monticola). The values of c _a-c _i were inferred from stable carbon isotope ratio (¹³C) of tree ring holocellulose adjusted for the 0.6–2.6 difference between holocellulose and whole sapwood. The cellulose extraction removed contaminants deposited in the tree ring after it formed and the adjustment corrected for the enrichment of cellulose relative to whole tissue. The whole sapwood values were then adjusted for bublished estimates of past atmospheric ¹³CO₂ and CO₂ concentrations. To avoid confounding tree age with CO₂, cellulose deposited by saplings in the 1980s was compared to cellulose deposited in the inner rings of nature trees when the mature trees were saplings, between 1910–1929 and 1941–1970; thus saplings were compared to saplings. In a separate analysis, the juvenile effect, which describes the tendency for ¹³C to increase in the first decades of a tree's life, was quantified independent of source CO₂ effects. This study provides evidence that conifers have undergone adjustments in the intercellular CO₂ concentration that have maintained c _a-c _i constant. Based on these results and others, we suggest that c _a-c _i, which has also been referred to as the intrinsic water-use efficiency, should be considered a homeostatic gas-exchange set point for these conifer species.     

Climate-related variability in carbon and oxygen stable isotopes among populations of Aleppo pine grown in common-garden tests

The Aleppo pine (Pinus halepensis Mill.) is found in the Mediterranean under a broad range of moisture and thermal conditions. Differences in severity and duration of water stress among native habitats may act as selective forces shaping the populations’ genetic make-up in terms of contrasting drought strategies. We hypothesised that these strategies should translate into intraspecific variation in carbon isotope composition (δ¹³C, surrogate of intrinsic water-use efficiency, WUE_i) of wood holocellulose, and such variation might be linked to changes in oxygen isotope composition (δ¹⁸O, proxy of stomatal conductance) and to some climatic features at origin. Thus, we evaluated δ¹³C, δ¹⁸O, growth and survival for 25 Aleppo pine populations covering its geographic range and grown in two common-garden tests. We found intraspecific variability for δ¹³C and growth, with high-WUE_i populations (which showed ¹⁸O-enriched holocellulose) having low growth. These results suggest stomatal regulation as common control for δ¹³C and productivity. We also detected sizeable relationships between δ¹³C and climate factors related to the magnitude and timing of drought such as the ratio of summer to annual rainfall. The main climate variable associated with δ¹⁸O was minimum temperature, but only in the coldest trial, suggesting differences in growth rhythms among sources. Overall, slow growing populations from highly-seasonal dry areas of the western Mediterranean exhibited a conservative water-use, as opposed to fast growing sources from the northernmost distribution range. The particular behaviour of the Mediterranean Aleppo pine as compared with other conifers demonstrates different selective roles of climate variables in determining intraspecific fitness.     

Using multiple trait associations to define hydraulic functional types in plant communities of south-western Australia

Assessing the hydrological imbalance and associated land degradation issues facing much of southern Australia and other parts of the world requires a better understanding of the defining features of ecosystem water use and the design of sustainable agroecosystems. Thus, by grouping species with similar water-use strategies into 'hydraulic functional types' (HFTs), we investigated the characteristics of water use in species-rich plant communities of south-western Australia. HFTs were determined using multiple-trait associations between morphological and physiological traits relating to water transport, water-use efficiency and response to water deficit. Sixteen traits were assessed from a subset of 21 species from three plant communities located along a topographically determined soil- and water-availability gradient. Multivariate analyses showed that trait variation was least at sites with shallower soils and putatively lower water availability, suggesting a convergence of water-use strategies at sites where plants are exposed to large seasonal water deficits. Stem hydraulic parameters, including stem-specific hydraulic conductivity, conduit diameter and maximum percentage embolism, were positively correlated, indicating the generality that larger conduit diameter permits greater hydraulic efficiency and is associated with greater seasonal reductions in hydraulic conductivity in this ecosystem. Wood density was not correlated with these traits, but closely associated with species' ability to withstand more negative water potentials during summer. Long-term integrated water-use efficiency was lower in shallow-rooted species that exhibited more negative summer water potentials. Specific leaf area and minimum leaf water potential were correlated with a number of separate traits, and appear to represent key axes of trait variation that describe the water-use strategies of different HFTs along the topographic gradient. Five HFTs were classified using a resemblance analysis according to combinations of traits that pertain to different water-use strategies among species; year-round active tree, year-round active shrub, hemiparasite, drought-suppressed broad-leaved shrub and drought-suppressed narrow-leaved shrub.     

Comparisons of carbon isotope discrimination in populations of aridland plant species differing in lifespan

Summary Carbon isotope discrimination () was compared between populations of dominant perennial plant species, differing in life expectancy, in two deserts with contrasting vegetation types. In both deserts, plants of the shorter-lived species showed significantly higher and greater intrapopulation variance in this character compared to the long-lived species. These results indicate underlying differences in gas-exchange physiology, and suggest a positive correlation between water-use efficiency and lifespan in desert plants. Differences in variance for this character may reflect greater microenvironmental variation experienced by shorter-lived plants and/or different forms of selection acting on water-use traits. Spatial distributions were significantly clustered for the shorter-lived species and significantly uniform for the long-lived species, indicating that competition has been important in the development of the long-lived populations. The long-lived Larrea tridentata showed a significant, negative correlation between and Thiessen polygon area, suggesting a positive relationship between water-use efficiency and longevity within this species. This relationship was weakly supported in the other warm desert species, Encelia farinosa, but was not observed within populations of the cold desert species, Gutierrezia microcephala and Coleogyne ramosissima. These results suggest that reflects key aspects of plant metabolism related to lifespan; these differences may ultimately influence interactions among desert plants and the structure of desert plant communities.     

Tree species from contrasting hydrological niches show divergent growth and water-use efficiency

Tree species occupy different hydrological niches and climate warming may affect tree performance in those niches through increased drought stress. However, the effects of climate warming on growth, carbon and water fluxes would differ depending on species’ hydrological niche. Species from wet sites should show a lower growth dependence on precipitation and also lower intrinsic water-use efficiency (WUEi), as compared with species from dry sites which should improve more the WUEi. We test these ideas by comparing radial-growth rates (basal-area increment), climate- and drought-growth associations and WUEi of hackberry (Celtis australis) vs. Aleppo pine (Pinus halepensis) and maritime pine (Pinus pinaster) in two Mediterranean sites located in Spain. Species are subjected to similar regional climate conditions in each site but occupy contrasting local hydrological niches (hackberry in wet sites and pines in dry sites). Climate is warming in both study sites and drought-avoiding pines are responding by showing higher growth rates and improved WUEi. We also found a similar growth dependency on winter-spring precipitation and summer drought of all species and sites and comparable WUEi values and trends, excepting in hackberry from southern Spain which grew more, and showed a higher growth resistance to drought and lower and more stable WUEi values. Variables inferred from tree rings as growth rates and WUEi allow characterizing the hydrological niche of tree species, which may be contingent on site conditions and climate warming.     

Leaf carbon isotope discrimination and vegetative responses of Dryas octopetala to temperature and water manipulations in a High Arctic polar semi-desert,Svalbard

Integrative ecophysiological and vegetative responses of Dryas octopetala were measured in response to field perturbations of temperature, precipitation and their interactions in a polar semi-desert in Svalbard, Norway (79°N, 12°E). Leaf carbon isotope discrimination (), total leaf nitrogen concentration and leaf development were determined for photosynthetic leaves collected during the last week of August 1991, after one season of manipulations. Individual leaf weight and the total mass of leaf tissue were significantly lower when water was added, irrespective of temperature regime. Leaf carbon isotope discrimination and estimated long-term c _i/c _avalues (the ratio of CO₂ concentration in leaf intercellular spaces to that in the atmosphere) were significantly higher under all three field manipulation treatments, and was significantly reduced when Dryas was grown under drought conditions in a related greenhouse study. Nitrogen concentrations of plants from the field experiment were significantly lower under warmed conditions regardless of water regime. Our results indicate that changes in environmental conditions in high arctic settings will result in alterations of Dryas leaf gas exchange, as expressed by increases in carbon isotope discrimination, which may be accompanied by shifts in leaf nitrogen content and leaf biomass.     

Physiological homeostasis for ecological success of Typha (Typha domingensis Pers.) populations in saline soils

The natural capacity of plants to endure salt stress is largely regulated by multifaceted structural and physio-biochemical modulations. Salt toxicity endurance mechanism of six ecotypes of Typha domingensis Pers. was evaluated by analyzing photosynthesis, ionic homeostasis, and stomatal physiology under different levels of salinity (0, 100, 200 and 300 mM NaCl). Typha populations were collected across different areas of Punjab, an eastern province in Pakistan. All studied attributes among ecotypes presented differential changes as compared to control. Different salt treatments not only affected gas exchange attributes but also shown significant modifications in stomatal anatomical changes. As compared to control, net photosynthetic rate, transpiration rate, total chlorophyll contents and carotenoids were increased by 111%, 64%, 103% and 171% respectively, in Sahianwala ecotype among all other ecotypes. Similarly, maximum water use efficiency (WUE), sub stomatal CO₂ concentration, sodium (Na⁺) and chloride (Cl⁻) contents were observed in Sahianwala (191%, 93%, 168%, 158%) and Knotti (162%, 75%, 146%, 182%) respectively, as compared to the others ecotypes. Adaxial and abaxial stomatal areas remained stable in Sahianwala and Knotti. The highest abaxial stomatal density was observed in Gatwala ecotype (42 mm²) and maximum adaxial stomatal density was recorded in Sahianwala ecotype (43 mm²) at 300 mM NaCl salinity. The current study showed that Typha ecotypes responded varyingly to salinity in terms of photosynthesis attributes to avoid damages due to salinity. Overall, differential photosynthetic activity, WUE, and changes in stomatal attributes of Sahianwala and Knotti ecotypes contributed more prominently in tolerating salinity stress. Therefore, Typha domingensis is a potential species to be used to rehabilitate salt affected lands for agriculture and aquatic habitat.Supplementary InformationThe online version contains supplementary material available at 10.1007/s12298-021-00963-x.     

Leaf-twig carbon isotope ratio differences in photosynthetic-twig desert shrubs

Summary Carbon isotope ratios were determined for bulk tissues of both leaves and current season twigs of 29 species of Mohave Desert shrubs. Leaf and twig tissues were found to differ in their carbon isotope ratios, only in those species which had photosynthetic twigs. These data suggest that the twigs of these species operate at lower intercellular CO₂ values than leaves, an interpretation which is consistent with available gas-exchange data. An effect of microhabitat was also evident between the mean isotope ratios of leaves from wash versus slope habitats.This research has been supported by NSF Grant BSR-8410943 and by DOE Grant DE-FGO2-86ER60399.